ARTICLE On the identity of (Orosarcophaga) ornata (Townsend, 1927) (Diptera: Sarcophagidae) and a new generic synonym

Cátia Antunes de Mello-Patiu¹² & Paula Raile Riccardi¹³

¹ Universidade Federal do Rio de Janeiro (UFRJ), Museu Nacional (MN), Departamento de Entomologia. Rio de Janeiro, RJ, Brasil. ² ORCID: http://orcid.org/0000-0002-0162-0952. E‑mail: [email protected] ³ ORCID: http://orcid.org/0000-0003-4850-7524. E‑mail: [email protected]

Abstract. The revision of the sarcophagid species Lepidodexia (Orosarcophaga) ornata (Townsend, 1927) reveals its identity and a generic synonym. The male terminalia of this species was studied in detail for the first time and revealed its congeneric affinities with Oxyvinia Dodge, 1966. Consequently, the generic status of Orosarcophaga Townsend, 1927 was revalidated, including Oxyvinia Dodge, 1966 as its junior synonym, and O. ornata was corroborated as a valid species and name. Moreover, we provide an updated generic diagnosis of Orosarcophaga, a list of the eleven species of the with their distribution, and a detailed diagnosis of O. ornata.

Keywords. Flesh ; Morphology; Sarcophagine; .

INTRODUCTION Orodexia and Orosarcophaga due to the need for a revision of Lepidodexia [s.l.], with the decision The genus Lepidodexia sensu Pape (1996) being made then by the first reviser. is composed of 32 subgenera, of which 11 are Townsend (1927: 342) described Orosarco- monotypic (e.g., Orosarcophaga), and it is rec- phaga ornata based on one male and one female ognized mainly by some features present in the specimens, showing only general coloration char- male terminalia (Pape, 1996; Buenaventura & acteristics. Later, Townsend (1938: 47) presented a Pape, 2018). For some Lepidodexia species, this more complete description of the genus and spe- structure was never represented in the literature, cies and designated the male specimen as a “ho- leading to some doubts about the generic place- lotype” [correctly, it should be lectotype], but the ment of such taxa, as is the case of Lepidodexia author did not provide any aspect of the termina- (Orosarcophaga) ornata. lia or illustrations. This type specimen had never The species currently recognized as Lepidodexia been studied since. Therefore, the male terminalia (Orosarcophaga) ornata (Townsend, 1927) was of the lectotype of O. ornata was studied in detail originally described in the monotypic genus for the first time herein, along with other speci- Orosarcophaga Townsend, 1927. Later, this genus mens recently collected in Brazil. was considered a junior synonym of Lepidodexia by Pape (1996), who gave a subgeneric status to Orosarcophaga, keeping only its type species. MATERIAL AND METHODS The same action was proposed by Pape (1996) for Orodexia Townsend, 1927 and Orobrachycoma The lectotype of Orosarcophaga ornata Townsend, 1927, which resulted in the second- housed at the National Museum of Natural History ary homonym of Lepidodexia (Orosarcophaga) (USNM), Smithsonian Institution, Washington, was ornata (Townsend, 1927), Lepidodexia (Orodexia) examined along with two additional specimens of ornata (Townsend, 1927), and Lepidodexia the Museu Nacional (MNRJ), Universidade Federal (Orosarcophaga) ornata (Townsend, 1927). Also, do Rio de Janeiro, Rio de Janeiro. For morpholog- Pape (1996) did not explain his criteria for this ical comparisons, seven specimens of six species taxonomic arrangement but stressed that no ac- of Oxyvinia and two specimens of Hallina retusa tion was taken to correct the homonymy between (Hall, 1933) from the MNRJ were examined, as well

Pap. Avulsos Zool., 2021; v.61: e20216135 ISSN On-Line: 1807-0205 http://doi.org/10.11606/1807-0205/2021.61.35 ISSN Printed: 0031-1049 http://www.revistas.usp.br/paz ISNI: 0000-0004-0384-1825 http://www.scielo.br/paz Edited by: Carlos José Einicker Lamas http://zoobank.org/7742C7F7-0017-49D0-8495-1921C3EC817F Received: 17/08/2020 Accepted: 15/01/2021 Published: 31/03/2021 Pap. Avulsos Zool., 2021; v.61: e20216135 Mello-Patiu, C.A. & Riccardi, P.R.: Identity of Lepidodexia (Orosarcophaga) ornata 2/6 as literature data on all known Oxyvinia species. We pro- base in profile; phallus with a distinct hinge between vide additional information on the examined material basi- and distiphallus; paraphallus bent in its proximal between brackets. third; well-developed vesica with proximal section (ves- Pictures were taken with the software LAS coupled to ical arm-shaped lever sensu Buenaventura & Pape, 2018) a stereomicroscope and assembled in software Helicon covering the remaining parts and distal section bifid; Focus 6. After the abdomen removal, the male termina- juxta attached to the paraphallus with a smooth surface, lia were treated with 10% KOH at 40 for 25 min, then often with a sclerotized distal portion and a membra- ℃ rinsed in 1% acetic acid and transferred to a microvial with nous proximal portion; median stylus with a basal pair of glycerin. Drawings were made using a camara lucida cou- elongated processes (capitis sensu Buenaventura & Pape, pled to a microscope, then edited using Adobe Illustrator 2018); robust lateral styli with an elongated base that ex- CS software. The external morphology nomenclature fol- tends laterally towards the paraphallus wall. lows Cumming & Wood (2017) and, for the distiphallus el- ements, Mello-Patiu & Pape (2000), except “median stylus” Distribution: Neotropical. and “paraphallus”, which are under the concepts by Giroux et al. (2010) and Whitmore et al. (2013), respectively. Comments: No females were available; therefore, they were not included in the generic diagnosis.

RESULTS Included species

After a detailed morphological study of Orosarcophaga angolensis (Hall) 1937: 374 (). Type locali- ornata, it is clear to state that this species indeed shares ty: Chile, La Araucanía, Angol. Distribution: Chile (La diagnostic features with members of the genus Oxyvinia Araucanía, Tarapacá). Ref. Lopes (1969); Pape (1996). Dodge, 1966, as those elucidated by Dodge (1966), Lopes New combination (1982), Pape (1996), and Buenaventura & Pape (2018). In excisa (Lopes) 1950: 361 (). Brazil, Rio consequence, we revalidate Orosarcophaga as a valid ge- de Janeiro, Guanabara, Grajaú. Distribution: Peru, neric name. Moreover, O. ornata does not resemble any Argentina, Brazil (Mato Grosso, Rio de Janeiro). Ref. other species currently allocated in Oxyvinia, and it is cor- Lopes (1969); Pape (1996); Dufek et al. (2020). New roborated as a valid species as well. combination Ten species have been recognized in Oxyvinia, but grata (Lopes) 1953: 44 (Dexosarcophaga). Type locality: Pape (1996) included eleven species in his catalog due Ecuador, Azuay, Cuenca. Distribution: Ecuador, Peru. to the inclusion of O. retusa (Hall, 1933). In the work by Ref. Lopes (1969); Pape (1996). New combination Pape (1996), this species was cataloged twice, as both grisea (Lopes) 1982: 290 (Oxyvinia). Type locality: Peru, Hallina (page 234) and Oxyvinia (page 272). Lopes (1975) Junin, San Ramon, Estancia Naranjal. Distribution: reviewed the species described by David Hall from Peru. Ref. Pape (1996). New combination Panama and furnished its redescription as Hallina retusa. ornata Townsend, 1927: 342 (Orosarcophaga). Type local- Our examination of the specimens deposited at the MNRJ ity: Brazil, São Paulo, Itaquaquecetuba. Distribution: also showed that H. retusa does not really resemble the Brazil (Rio de Janeiro, São Paulo). Ref. Lopes (1969); Oxyvinia species, and we did not include it in our checklist. Pape (1996); additional material. New status Below, we provide an updated generic diagnosis of panamensis (Lopes) 1988: 132 (Oxyvinia). Type locality: Orosarcophaga, including features of the male termina- Panama, Chiriqui, Potrecillos. Distribution: Panama. lia, a list of the eleven species of the genus with their dis- Ref. Pape (1996). New combination tribution, and a detailed diagnosis of O. ornata. piliventris (Dodge) 1966: 692 (Oxyvinia). Type locality: Venezuela, Cerro Avila, nr [near] Caracas. Distribution: Venezuela. Ref. Lopes (1969); Pape (1996). New Genus Orosarcophaga Townsend combination uraricoera (Tibana & Lopes) 1990: 669 (Oxyvinia). Type Orosarcophaga Townsend, 1927: 231. Type species: locality: Brazil, Roraima, Rio Uraricoera, Maraca I. Orosarcophaga ornata Townsend, 1927, by original Distribution: Brazil (Roraima). Ref. Pape (1996). New designation. New status combination Oxyvinia Dodge, 1966: 692. Type species: Oxyvinia vittata (Lopes) 1982: 288 (Oxyvinia). Type locality: Bolivia, piliventris Dodge, 1966, by original designation. New El Limbo. Distribution: Bolivia. Ref. Pape (1996). New synonym combination wicharti (Lopes) 1953: 46 (Dexosarcophaga). Type local- Generic diagnosis: Rows of almost parallel frontal setae; ity: Brazil, Rio de Janeiro, Itatiaia. Distribution: Brazil parafacial plate with setulae only; four dorsocentral post- (Rio de Janeiro), Colombia. Ref. Lopes (1969); Pape sutural setae; postalar wall setose; vein R₁ bare; ctenid- (1996), Buenaventura et al. (2020). New combination ium on mid femur; male ST 5 with cushion-like protu- xanthophora (Schiner) 1868: 313 (Sarcophaga). Type berances in the inner margins of the cleft (Fig. 2B; arrow) locality: Venezuela [“Süd-Amerika”]. Distribution: covered by setae or spines; reddish or yellowish-brown Ecuador, Peru, Venezuela, Brazil (Paraná). Ref. Lopes male terminalia; apically curved pregonite relative to the (1969); Pape (1996). New combination Mello-Patiu, C.A. & Riccardi, P.R.: Identity of Lepidodexia (Orosarcophaga) ornata Pap. Avulsos Zool., 2021; v.61: e20216135 3/6

Orosarcophaga ornata Townsend, 1927 New status pedicel apex level; reclinate orbital seta present, procli- (Figs. 1‑2) nate orbital setae absent; outer vertical setae 0.3x inner vertical; black ocellar triangle, ocellar setae as developed Type material: Lectotype male (USNM) [designated as frontals; postocular area with yellowish-gray pruinos- by Townsend 1938: 47, destroyed]. Brazil, São Paulo, ity; gena with golden pruinosity and black setae; black Itaquaquecetuba (Fig. 1C). The lectotype label (Fig. 1C) postgena with slightly silvery pruinosity, black setae and does not include information on the locality, which was some few whitish setae close to occiput; face with gold- retrieved from Lopes (1969). en pruinosity; black facial ridge with short setulae in the inferior half; black antenna, first flagellomere with gray Additional material: 2 Brazil, Rio de Janeiro, Itatiaia, pruinosity and about 2x longer than pedicel; arista long ♂♂ Parque Nacional de Itatiaia, trilha [= trail] Rui Braga, plumose on basal 2/3; blackish palpus. 1,198 m, 22°26 09.1 S, 44°37 31.2 W, 12‑18.iii.2017, Van ′ ″ ′ ″ Someren – shrimp [bait], Nihei et al. leg. [MNRJ] Thorax (Figs. 1A‑B): Black with intense golden pruinosity on mesonotum and scutellum, postpronotum, notopleu- Diagnosis: Head and thorax with intense golden pru- ron, anepisternum, anepimeron, and katepisternum. inosity, black abdomen with silvery-gray pruinosity; Chaetotaxy: acrostichals 2‑3 + 1, dorsocentrals 4 + 4 (2 syntergosternite 7 + 8 with blackish-brown anterior half longer posteriormost), intra-alars 1 + 2, supra-alars 2 + 3, and yellowish-brown posterior half; yellowish-brown ep- postpronotals 3, notopleurals 4; katepisternals 3 setae andrium, cerci, and phallus; cercal prong with acute and almost in a straight line; setulose postalar wall; postalar curved forward apex, in profile; pregonite with apical callus with 2 setae; meral setae 10‑11; bare proepister- half curved and spatulate; distally dentated postgonite; num; setulose prosternum; scutellum with a pair of basal, juxta broad and not detached from the paraphallus, dis- lateral, and subapical setae (the lateral one shorter and tal margin with a pair of teeth in the middle accompa- near the subapical), a pair of preapical discal, and a pair nied by 2‑3 teeth on each side; vesica with the surface of apical setae. Wing. Hyaline, with dark-brown veins; covered with microtrichiae, with two hook-like projec- yellow basicosta and black tegula; bare vein R₁; vein R₄₊₅ tions, in profile; broad median stylus, with one basal pair with setulae dorsally on almost the entire distance to of elongated processes; long and robust lateral styli. crossvein r‑m; cell r₄₊₅ open at wing margin; third cos- tal sector bare ventrally; costal spine not differentiated. Male: Length: 7‑9 mm (n = 3). Legs. Blackish-brown with silvery-gray pruinosity, yellow- ish-brown pulvilli; mid femur with a row of 3 median an- Head (Figs. 1A‑B): Parafacial and fronto-orbital plates with terior setae, a row of anteroventral setae, 2 preapical pos- intense golden pruinosity; parafacial plate with a row of terior setae, a row of posteroventral setae with an apical setulae close to the eye; frons about 0.30x head width at ctenidium of 8‑9 modified setae; mid tibia with 1 median ocellar triangle level; entirely blackish frontal vitta; rows anterodorsal, 1 apical dorsal setae, 2 basal, 1 median, and of parallel frontal setae except two slightly divergent an- 1 apical posterodorsal setae; hind femur with a row of an- teriormost, 9‑10 well-developed frontal setae reaching terodorsal setae, a median row of anterior setae, a row of

Figure 1. Orosarcophaga ornata lectotype. (A) Habitus (Scale bar: 1 cm). (B) Dorsal view. (C) Label. Pap. Avulsos Zool., 2021; v.61: e20216135 Mello-Patiu, C.A. & Riccardi, P.R.: Identity of Lepidodexia (Orosarcophaga) ornata 4/6

Figure 2. Orosarcophaga ornata postabdomen. (A) Male postabdomen in profile. (B) Fifth sternite; arrow indicates a cushion-like protuberance. (C) Phallic complex in profile (lectotype). (D) Phallus in ventral view (lectotype); arrow indicates an elongated process on the medial stylus. Scale bar: 1 mm. Abbreviations: ap pg, postgonal apodeme; bas, basiphallus; cer, cercus; ep, epandrium; hyp, hypandrium; ju, juxta; phal, phalapodeme; pog, postgonite; preg, pregonite; S 7 + 8, syn- tegosternite 7 + 8; st l, lateral stylus; st m, medial stylus; sur, surstylus; ves, vesica. Mello-Patiu, C.A. & Riccardi, P.R.: Identity of Lepidodexia (Orosarcophaga) ornata Pap. Avulsos Zool., 2021; v.61: e20216135 5/6 anteroventral setae, 1 preapical dorsal, and 1 preapical phylogenetic hypothesis of proposed by posterior setae; hind tibia with 1 basal, 1 median, and 1 Buenaventura & Pape (2018), three species of the for- apical setae in the same position on the anterodorsal and mer Oxyvinia were nested with Dexosarcophaga species; posterodorsal margins, and 1 median anterior seta; hind however, Orosarcophaga ornata was not included in the coxa and trochanter with normal setae. taxonomic sample. Riccardi & Mello-Patiu (2020) nested Orosarcophaga with Townsend instead of Abdomen (Figs. 1A‑B, 2B): Black with silvery-gray pru- Dexosarcophaga Townsend, but the taxon sampling of inosity; T3 with 1 lateral marginal and no median mar- Sarcophaginae except Lepidodexia was limited. Despite ginal setae; T4 with 3 lateral marginal and 1 median mar- the differences aforementioned, both phylogenies indi- ginal setae; T5 with a complete row of marginals (ca. 20); cate that Orosarcophaga (including Oxyvinia) does not rectangular ST2‑4, ST2‑3 with numerous long hair-like belong to Lepidodexia. pale setae and some black ones; ST3‑4 with a patch of The male terminalia elements of sarcophagids are black setae on the middle of the posterior margin; black- often the most reliable morphological features to es- ish ST5 with a cushion-like elongated process in the inner tablish phylogenetic relationships (Giroux et al., 2010; margins, covered by spinous setae (Fig. 2B). Buenaventura & Pape, 2018). In this sense, the most con- spicuous characteristic gathering the members of the Terminalia (Figs. 2A, 2C‑D): Syntergosternite 7 + 8 with former Oxyvinia into Orosarcophaga is the median stylus blackish-brown anterior half and yellowish-brown pos- with a peculiar basal pair of elongated processes, which terior half, 4‑5 pairs of well-developed marginal setae; seems to be a unique feature within Sarcophaginae yellowish-brown epandrium, cerci, and phallus; epan- (Fig. 2D; arrow). Additionally, the type species of drium with sparse setae, longer on the dorsal surface Orosarcophaga also share with O. vittata and O. xantho- (Fig. 2A); clavate surstylus with long setae on the apical phora the presence of a dentate margin on the apex of half (Fig. 2A); broad cercal prongs, parallel but separat- juxta. This evidence, along with the remaining features ed, in posterior view, with acute, blackish, and curved discussed above, supports our taxonomic decision. forward apex, in profile (Fig. 2A); long pregonite, api- cal half curved and spatulate, and postgonite as long as the pregonite, with a long median setae and distally ACKNOWLEDGMENTS dentate (Fig. 2C); separated basi- and distiphallus, short basiphallus, about 0.5x the paraphallus length (Fig. 2C); We thank Norman E. Woodley (USNM) for the loan of broad juxta not detached from the paraphallus, distal the lectotype. C.A.M.P. is grateful to FAPERJ for its finan- margin with a pair of teeth in the middle accompanied cial support (Proc. E‑26/200.078/2019) and CNPq for the by 2‑3 teeth on each side (Figs. 2C‑D); pilose vesica with Research fellowship (308951/2018‑2). P.R.R. is support- two hook-like projections, in profile, which are an odd ed by a postdoctoral fellowship from the CAPES/CNPQ upper lobe covering a bifid lower lobe, in ventral view Acordo Program – PROTAX II – Project 440482/2015‑1 (Figs. 2C‑D); broad median stylus with one basal pair of (Proc. 176040/2018‑00 and 333912/2019‑00). elongated processes (Fig. 2D; arrow); long and robust lat- eral styli (Fig. 2D). AUTHORS’ CONTRIBUTIONS Remarks: The lectotype was on loan at the MNRJ and, unfortunately, it was lost during the fire which destroyed C.A.M.P.: contribution statement – this author was the the MNRJ Entomological Collection (Duarte, 2019). We major contributor of the manuscript, provided the back- observed that there are individual differences in the ground history of Orosarcophaga, the updated generic number of teeth in the juxtal lateral dentation among diagnosis, the list of species, and analyzed the remaining the specimens examined (2 or 3 teeth on each side of the species of the genus. P.R.R.: contribution statement – this median teeth pair). author provided insights into all sections of the manu- script, the images and illustrations, participated actively in the material and methods section, the written charac- DISCUSSION terization of Orosarcophaga ornata, and the discussion.

Despite the presence of setae on the postalar wall, Orosarcophaga ornata does not possess the vesica REFERENCES C‑shaped with a convex distal section, juxta angled rel- ative to the phallic tube, developed bacilliform scler- Buenaventura, E. & Pape, T. 2018. Phylogeny, evolution and male terminalia ite, nor a spinose lobe above the vesica, all diagnostic functionality of Sarcophaginae (Diptera: Sarcophagidae). Zoological characteristics of the genus Lepidodexia (Pape, 1996; Journal of the Linnean Society, 183(4): 808‑906. Buenaventura & Pape, 2018; Riccardi & Mello-Patiu, Buenaventura, E.; Valverde-Castro, C. & Wolff, M. 2020. New carrion-visiting 2020). Also, the result of an ongoing morphological flesh (Diptera: Sarcophagidae) from tropical dry forests of Colombia phylogeny of Lepidodexia shows O. ornata nested with and their phylogenetic affinities. Acta Tropica, 213(2021): 1‑14. DOI O. xanthophora, separated from all the remaining spe- Cumming, J.M. & Wood, D.M. 2017. Adult morphology and terminology. In: cies of Lepidodexia (Riccardi & Mello-Patiu, 2020). In the Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.). Manual of Afrotropical Diptera. Pap. Avulsos Zool., 2021; v.61: e20216135 Mello-Patiu, C.A. & Riccardi, P.R.: Identity of Lepidodexia (Orosarcophaga) ornata 6/6

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